Foraging: The Game of Give and Take

Bolaños, Victoria

Abstract
Food provides nutrients and energy to primates. I studied behavioral activities of wild

Cebus capucinus, Ateles geoffroyi, and Alouatta palliata and compared it to the type of food they

ate. Fruit provides higher energy benefits to primates than leaves. It was predicted, the primates

would eat more fruit and would move and travel more than any other behavioral activity. The

hypothesis was based on the Optimal Foraging Theory that bases foraging on costs, rewards, and

constraints. This study was conducted at La Selva Biological Station in Puerto Viejo de

Sarapiqui, Costa Rica. The study site consisted of primary forest, secondary forest, select logged

forests, early successional pasture, and abandoned plantations. The Scan Method was used to

collect data in 2 minute scan intervals (Campbell et al., 2011) (Altmann, 1974). A. palliata ate

mostly leaves and spent most of their time resting. C. capucinus spent most of their time moving

and their food source was a non-leaf part of a palm tree. A. geoffroyi only ate leaves and spent

most of their time moving. My hypothesis on A. palliata was rejected. There were limited data

collected on C. capucinus and A. geoffroyi to draw a conclusion. There may be additional factors

that influence foraging behavior in addition to the Optimal Foraging Theory. Foraging is a

complex concept that requires understanding multiple influential factors.

Introduction
The Optimal Foraging Theory states foraging is based on three concepts: costs, rewards,

and constraints. The amount of time and energy to forage for food is the “cost”. The “rewards”

are the nutritional values gained from the food. The “constraints” are factors that influence the

cost and rewards (as cited in Dzul-Cauich et al., 2018). Fruits have higher nutritional benefits

and provide higher energy levels to primates than leaves. Nutritional contents in food that

influence the “rewards” are protein, fiber, carbohydrates, etc. (Garber 1987). Cebus capucinus,
Ateles geoffroyi, and Alouatta palliata all consume fruit when available (Beader et al., 2011).

Primates have the ability to adapt their diet based on their surroundings.

Capuchin Monkeys

C. capucinus spend about 75% of their day foraging. C. capucinus eat fruit, seeds, bugs,

and small prey (Beader et al., 2011). They can survive on a diet of just bromeliads or seeds

during food shortages. They do eat small prey such as birds, lizards, squirrels, and coatis. In C.

capucinus groups, if an individual captures a prey, they can eat it with minimal harassment from

their group (Beader et al., 2011). C. capucinus are known to raid plantations and farms, mainly

for oranges, corn, or anything available (Britannica, 2017). C. capucinus are well known for their

use of tools in the wild, such as a club or leaves, to help with foraging (Beader et al., 2011).

Howler Monkeys

A. palliata enjoy eating fruits and leaves. In primary forests, the A. palliata diet consists

of 50% leaves, 30% fruits, and 15% flower (de Thoisy & Richard-Hansen, 1997). Due to

seasonal availability of fruits, they eat leaves for most of the year. They particularly enjoy fruits

and leaves from fig trees. Leaves on plants will contain toxins to help deter foraging from

animals, therefore, A. palliata need to eat various types of leaves to fulfill their dietary needs

(Amazon Aid Foundation).

Spider Monkeys

A. geoffroyi consume decaying wood and drink water or consume soil at mineral licks. A.

geoffroyi enjoy eating lipid rich fruits (Beader et al., 2011). Their diet consists of about

83%-90% of fruit. Their remaining diet consists of flowers, leaves, honey, nuts, insects, bird

eggs, sap, and bark (Monkey, Spider). The dominant female will determine the route for foraging

as well as the time of day for foraging (Monkey, Spider). Females tend to have a greater variety

of food in their diet than males (Spider Monkey, Lamar University). A. geoffroyi are able to

swallow fruits whole, including the seeds (New World Encyclopedia).

 Due to the low energy levels leaves provide, A. palliata rest about 80% of the day and are

known for being least active compared to other primates (Howler Monkey, New World

Encyclopedia). The high fruit diets of C. capucinus and A. geoffroyi provide high “rewards” in

the form of energy to them so they are able to be active. The high leaf diet of A. palliate provides

low “rewards” therefore, they must use their energy sparingly and consciously.

I predict all three primates will consume more fruit than leaves due to the higher rewards

fruits provide than leaves. I also predict, all species of primates studied will spend more time

moving and travelling than feeding and resting due to the high energy rewards provided in fruits.

Materials and Methods

Study Area

The primates were studied at La Selva Biological Station, located at the confluence of the

Sarapiqui and Puerto Viejo rivers (McDade & Hartshorn, 1994) (Fig. 1). La Selva is located in

the North West region of Costa Rica in Puerto Viejo de Sarapiqui in the Heredia Province. La

Selva has been owned by the Organization for Tropical Studies since 1968 (La Selva Research

Station). The biology station sits on 1,536 hectares. The extended areas of La Selva include La

Guaraia Annex, East Bank, La Flaminea, and an area extending from the East Bank to the Puerto

Viejo-Horquetas Highway. La Selva’s land consists of primary forest, secondary forest, select

logged forests, early successional pasture, and abandoned plantations. Primary forests are forests

that have not been disturbed and have mature trees. Secondary forests are regenerating forests

that have experienced mass loss by natural or manmade causes. The vegetation can be unstable

as it is experiencing successional stages (Forest). The secondary forest areas range from

10 years - 47 years. There is about 157 inches of rainfall annually at La Selva. The dry season is

rarely long or severe. The daily temperature averages from 6° C to 12°C (McDade & Hartshorn,

1994). There are over 2,077 species of plants (La Selva Research Station).
Figure 1. A trial map and land use of La Selva Biological Station.

(Organization for Tropical Studies)

(La Selva Biological Station GIS Lab, 2018)

 Study Subjects

Capuchin Monkeys
C. capucinus live in the region of Honduras through the northwestern coast of Ecuador.

They are native to South America. (Jack 2011). Male C. capucinus weigh 3.7-3.9 kg while

females weigh 2.6-2.7 kg (Jack 2011). Their forelimbs and hindlimbs are equally proportional

(Jack 2011). They have short fingers and all digits move independently. They also have “pseudo-

opposable thumbs” (Jack 2011). C. capucinus do not have tufts on the top of their heads (Jack

2011). They have terrestrial locomotor patterns due to their equal limbs. Since their tails are full

of fur, they are unable to maintain their full body weight on their tail. Their tail provides support

and balance while they are foraging. Their tail acts like an anchor to help them balance on tree

limbs (Jack 2011).

The average group size of C. capucinus is 16.4. Males tend to be the dominate sex. Alpha

females do exist but they are viewed as below the alpha male (Jack 2011). The alpha female can

dominate subordinate males (Jack 2011). “Females direct more threats towards males than they

receive from them” (Jack 2011).

Howler Monkeys

A. palliata are found in moist tropical and deciduous forests. They are found between

Northern Bolivia to Southern Mexico and the Yucatan Peninsula (Di Fiore et al. 2011).

A. palliata tails and limbs are considered elongated compared to their trunk length. They have a

reduced thumb while their remaining digits are elongated. A. palliata have a muscular prehensile

tail that supports their full body weight when feeding and during locomotion (Di Fiore et al.

2011).

A. palliata females in new groups typically join because of an emigrating male. These

new females are unrelated to members of the group. Over time, the founding females will try to

recruit their own daughters as breeders. They also try to force emigration of the other daughters

in the group (Di Fiore et al. 2011).

Spider Monkeys

A. geoffroyi, are also found in moist tropical and deciduous forests between Northern

Bolivia to Southern Mexico and the Yucatan Peninsula (Di Fiore et al. 2011). Similar to

A. palliata, A. geoffroyi, have tails and limbs that are elongated compared to their trunk length.

They also have a reduced thumb while their remaining digits are elongated. A. geoffroyi have a

muscular prehensile tail that supports their full body weight when feeding and during locomotion

(Di Fiore et al. 2011).

A. geoffroyi live in multimale/multifemale communities (Di Fiore et al. 2011). They have

a fission-fusion group pattern as well. It is thought that a benefit of a fission-fusion society is the

group has an ability to adapt to predation (Di Fiore et al. 2011). One downside of this society

though, is a possibility of scarce food supplies. A. geoffroyi will forge together. Their social

system is sometimes described as "sex-segregated" because adult females and their offsprings

will forage and travel together while the adult males will range farther away (Di Fiore et al.

2011). Males tend to spend more time with other males than with females. There are often more

interactions of males together than mixed genders or female/female interactions. Adult males

tend to groom baby males more than female adult/female baby and mixed sex groups (Di Fiore et

al. 2011). Females tend to range separately from other females in their community. Males range

further and longer than females. Males patrol the boarders of their communities. They tend to be

aggressive in intercommunity interactions (Di Fiore et al. 2011).

Data Collection Method

Behavioral data on C. capucinus, A. geoffroyi, and A. palliata were collected from

May 9th – 19th, 2019. Data were collected over a total of 42 hours and 52 minutes over 26,535

meters on 11 trails in Primary, Secondary & Selected Logged Forests. There were not more than

2 consecutive days of no observations. To measure the rate of each behavior, the Scan Method

(Campbell et al., 2011) was used in 2 minute intervals (Altmann, 1974) during the hours of
5:00 AM – 5:10 PM. This allowed me to collect data for a large group as well as find the

percentage of time spent on each activity during the study period (Altmann, 1974). The first

activity seen by each individual was recorded (Jung et al., 2015). Only individuals within view

were recorded. The behaviors were placed in the following categories: resting, moving, traveling,

feeding, and other (social play, scratching, etc.) (Fig. 2). Any feeding was categorized as either

fruit, leaf, or other (wood, nuts, seeds, bugs, etc.) (Jung et al., 2015).

Figure 2. The Ethogram describing each type of recorded behavior.

Behavior Description
Resting No moving in a sitting, laying, or
standing position
Moving Locomotion within the same
tree (Jung et al., 2015).
Traveling Locomotion between trees
(Jung et al., 2015) or on the
ground
Feeding Food is in the hand or mouth,
chewing, and swallowing (Jung
et al., 2015)
Other e.g. Grooming, scratching,
urination, defecation, social
play, etc.

Results

There were a total of 182 scans in which A. palliata were in view during 6 hours

and 53 minutes of behavioral observations. There were 286 behavioral and feeding occurrences

observed on A. palliata (Fig. 3). All observations were in a Secondary & Selectively Logged

Forest area. The group sizes ranged from 3-7 individuals including babies, juveniles, and adults.

Resting occurred the most than any other behavioral activity, it accounted for 54% of the total

behavioral occurrences that was not feeding (Fig. 4). A. palliata was observed eating leaves 55%

of the time. I was unable to identify some of the food eaten which accounted for 40% of their

feeding behavior. Due to the lower “rewards” levels leaves provide nutritionally, it would be

expected that A. palliata would need to spend more time resting to not over exert themselves.
 A. geoffroyi were observed for 10 minutes in which 5 scans they were in view (Fig. 3) in

a Primary Forest. There were only 2 adults that were observed. Most of the observed behavior

was moving. Moving occurred 75% of their behavioral observations that was not feeding (Fig.

4). They ate leaves during the entire feeding observation. Both adults moved out of site. It is

surprising to see how active they were considering they were only observed eating leaves. Since

their observation time was short, it is possible they may have eaten high nutritional foods prior to

the observation. Longer observations would be needed to draw a conclusion.

There were a total of 6 scans in which C. capucinus were in view during 20 minutes of

behavioral observations. The group size consisted of 4 adults and 1 baby. One scan had 3

individuals in site. One scan had one induvial in view. The remaining scans had 2 adults within

view. During this time, there were 10 behavioral occurrences observed (Fig. 3) on C. capucinus

in a Secondary & Selectively Logged Forest. The food source they ate was from the palm tree

they were on that wasn’t the leaf. This accounted for 100% of all recorded feedings (Fig. 4).

Moving accounted for 67% of their behavioral activity not related to feeding. It is likely their

food source provided higher “rewards” levels than leaves allowing them to be active.

Figure 3. The total number of behavioral activity and food eaten occurrences.

A. Palliata Behavioral A. Palliata Food Source

Activity Occurrences
Number of Food Source Occurrences

140 35
Number of Observations

120 30
100 25
80 20
60 15
40 10
20 5
0 0
Resting Moving Traveling Other Fruit Leaf Other
Behavioral Activity Type of Food Consumed
 Figure 3 (Continued). The total number of behavioral activity and food eaten occurrences.

C. Capucinus Behavioral C. Capucinus Food Source

Activity Occurrence

Number of Food Source Occurrences

4.5 1.2
4
Number of Observations

1
3.5
3 0.8
2.5
0.6
2
1.5 0.4
1
0.2
0.5
0 0
Resting Moving Traveling Other
Behavioral Activity Type of Food Consumed

A. Geoffroyi Behavioral A. Geoffroyi Food Source

Activity Occurrence
Number of Food Source Occurrences

3.5 1.2
Number of Observations

3 1
2.5
0.8
2
0.6
1.5
0.4
1
0.5 0.2

0 0
Moving Traveling Leaf
Behavioral Activity Type of Food Consumed

Figure 4. The total percentage of food source eaten and each behavior recorded.

A. PALLIATA A. PALLIATA
Percentage of Food Source Eaten Percentage of Behavioral Activity
Other
Fruit 6%
5% Traveling
4%
Other
40%

Resting
54%

Leaf
55% Moving
36%
 Figure 4 (Continued). The total percentage of food source eaten and each behavior recorded.

A. GEOFFROYI A. GEOFFROYI
Percentage of Food Source Eaten Percentage of Behavioral Activity

Leaf Traveling
100% 25%

Moving
75%

C. CAPUCINUS C. CAPUCINUS
Percentage of Food Source Eaten Percentage of Behavioral Activity

Other
100% Traveling Resting
17% 16%

Moving
67%

Conclusion

I predicted the primates would eat mostly fruit and would move and travel most of

their time due to the high rewards fruits provide. My hypothesis was not supported by the

observations on A. palliata. They spent most of their time resting instead of moving and

traveling. Also, A. palliata ate more leaves than anything else, which rejects my hypothesis. This

does support the Optimal Foraging Theory, since they ate leaves that provided low energy

nutrients they needed to conserve their energy by limiting their activities.

 There were little behavioral data observed on C. capucinus and A. geoffroyi to draw a

definitive conclusion on my hypothesis. The data collected on C. capucinus showed they spent

the most amount of time moving. Also, the food C. capucinus were observed eating were not

leaves. It is possible the food source from the palm trees contained a slightly higher energy

nutrient than leaves which may have allowed them to be active. The data observed on

A. geoffroyi rejected my hypothesis, they only ate leaves and were highly active. Longer

behavioral observations are needed on C. capucinus and A. geoffroyi to draw a conclusion.

Despite my best efforts, locating C. capucinus and A. geoffroyi was the most challenging

aspect of this research. Due to the limited observation times for both species, a conclusion could

not be drawn on them. For follow up studies on this topic, more observational time will allow for

a better understanding of how food selection effects behavioral activities.

Discussion

In contrast to my prediction, A. palliata consumed mainly leaves thus needing to

rest more often. This data supports the Optimal Foraging Theory. The largest A. palliata group

size contained 7 individuals including 5 adults and 2 babies. Larger groups tend to have lower

foraging costs by having resource defense and by providing useful information regarding a site’s

food quality and size to members of the group (Garber 1987). As noted in Dzul-Cauich et al.

2018 research between fruit choices in A. pigra, A. geoffroyi, and bats in Southern Mexico, there

may be a relationship between A. pigra and the large tree, E. cyclocarpum. The tree seems to

provide an energy balance for the A. pigra by providing both food and a habitat for them to rest

and have social interactions with each other (Garber 1987). Searching for food can have a high

“cost” value for primates, especially for those that mainly eat leaves. To increase their

“rewards”, finding a location that provides both food and a safe shelter would be ideal. My study

showed A. palliata were able to find a tree that provided both food and safety for them to rest to
balance out their “costs” and “rewards”. These are possible reasons why A. palliata may have ate

leaves and rested the majority of the observations.

However, A. geoffroyi did not follow the model of the Optimal Foraging Theory since

they only ate leaves and were mostly active. This may be because there were only 2 adult A.

geoffroyis, so they had a higher foraging cost (Garber 1987). They did not have a defense

resource since their group size was small so they had to continue searching for food as quickly as

possible.

C. capucinus were active and although, C. capucinus were not seen eating fruits, they

did eat a non-leaf food source. It is possible this food source provided them higher nutritional

values, such as fiber, carbohydrates, protein, etc., than leaves which may have provided them

with higher “rewards”.

Future studies on the Optimal Foraging Theory can help increase the survival of animal

species by understanding their behavior and diet. According to Garber 1987, foraging is a

complex concept that may involve additional factors such as digestive morphology, resource

defense, group size, within group competition, or social organization to name a few. Additional

studies can help further understand foraging behavior.

Acknowledgements

I thank Salt Lake Community College and their Study Abroad Office, and Professor Melissa

Seaboch for coordinating this research. Thank you Professor Melissa Seaboch for the help and

support of this research and providing feedback. Thank you to La Selva Biological Center for

allowing this research to be conducted.

Conflict of Interest

I, declare there is no conflict of interest.

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